Cushman Foundation Special Publication No. 46, p.
393-402, 2018
Chapter 13
TAXONOMY, BIOSTRATIGRAPHY, AND PHYLOGENY OF OLIGOCENE
ACARININA
Bridget S. Wade1 and Morana Hernitz Kucenjak2
1
Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT, U.K.
Email: b.wade@ucl.ac.uk
INA-Industrija nafte d.d., Exploration and Production, Lovinciceva 4, 10 002 Zagreb, Croatia.
2
Email: morana.hernitz-kucenjak@ina.hr
ABSTRACT
The taxonomy, phylogeny and biostratigraphic
ranges of three Oligocene species of Acarinina are
discussed together with their synonymies. Acarinina
diversified in the Paleocene and Eocene and only a
INTRODUCTION
The muricate wall texture which is characterized
by multiple pustules on the test surface is present in
many early Paleogene planktonic foraminifera (e.g.,
Morozovelloides, Acarinina, Morozovella, Igorina,
Pearsonites). All these genera were abundant in tropical /
subtropical oceans of the early Paleogene. Stable isotope
data suggest that muricate forms dominated in the upper
part of the water column and have been inferred as
mixed-layer dwellers (e.g., Boersma and others, 1987;
Pearson and others, 1993; Wade and Kroon, 2002; Wade,
2004; Luciani and others, 2017a). Size fraction carbon
isotope analysis also suggests that the muricate forms
had a symbiotic relationship with algae (e.g., Pearson
and others, 1993; Norris, 1996; Kelly and others, 1998;
Wade and others, 2008; Luciani and others, 2017b).
The taxonomy and phylogeny of Paleocene
and Eocene Acarinina has been addressed in Olsson
and others (1999) and Berggren and others (2006),
respectively. The diversity of the acarininids decreased
through the Eocene, with 16 species recognized in the
latest early Eocene (~50 Ma), diminishing to 6 in the
few species remain in the OligoceneThe following
taxa are recognized as valid species: Acarinina
collactea (Finlay), Acarinina echinata (Bolli),
and Acarinina medizzai (Toumarkine and Bolli).
latest middle Eocene (~40 Ma; Berggren and others,
2006). The turnover of planktonic foraminifera at the
end of the Bartonian saw the demise of many of the
muricate forms with the extinction of Morozovelloides
and the large (>250 µm) acarininids (Wade, 2004;
Wade and others, 2012). Small acarininids continued
in the <125 µm size fractions through the latest Eocene
(Wade, 2004; Berggren and others, 2006; Agnini and
others, 2011) but it has generally been accepted that
all muricate forms were extinct prior to the Eocene
/ Oligocene boundary (Berggren and others, 2006).
However, our studies of Oligocene sediments from
multiple locations indicate that the acarininid lineage
continued through the Oligocene and perhaps into the
Miocene. These documented occurrences of muricate
forms in the Oligocene are significant and suggest that
this ancient and now extinct group continued for >10
myr later than previously documented. All these forms
are distinctive, small in size (<125 µm), and thought
to be in situ because they occur in the absence of other
small reworked species. Here we report and illustrate
Oligocene muricate planktonic foraminifera assigned to
Acarinina from the Pacific, Indian and Atlantic Oceans.
 Wade and Hernitz Kucenjak

We suggest that these taxa are not reworked but true

descendants of Eocene Acarinina and an extension of

Antarctic
tropical

tropical
the ancient muricate phylogeny.

(Sub)

(Sub)
The Oligocene acarininids are generally rare
Acarinina
and never dominate the assemblages like their Eocene
Cande & Kent (1995)

ancestors. It has therefore been difficult to constrain their

GPTS Age (Ma)

N Zones (K&S, 1983)

ranges and phylogeny. The Oligocene acarininids differ
E, O and M Zones

Huber & Quillévéré

Former P Zones
(BKSA, 1995) &
Epoch

from early Paleogene forms in that they are characterized

(WPBP, 2011)
by muricae which are blunter, shorter, and more widely
(2005)
dispersed. They differ from dipsidripellids by their

collactea

medizzai
echinata
normal perforate wall and more evenly distributed
muricae (rather than short hispid pustules). While the
large Eocene acarininids are well documented as having
been mixed-layer dwellers with photosymbionts, it is not
24
N4

? clear whether these small Oligocene forms shared that

a

ecological preference. The species level range-chart is

O7 presented in Figure 13.1.
25
In addition to the species discussed, Acarinina
AO4

inaequiconica Subbotina, 1960 was described from

26 P22 Oligocene sediments in the Ciscarpathian region of
LATE

Ukraine (Subbotina and others, 1960). Our new SEMs of

O6 the holotype reveal a small, low trochospiral specimen,
27 but the poor preservation has inhibited the determination
OLIGOCENE

of the taxonomic affinity of this form (see Chapter 20,

AO3

28 b O5 this volume).
P 21

a O4 SYSTEMATIC TAXONOMY
29
AO2

Order FORAMINIFERIDA d’Orbigny, 1826

30 P20 O3 Superfamily GLOBIGERINOIDEA
Carpenter, Parker, and Jones, 1862
EARLY

Family TRUNCOROTALOIDIDAE
31 P19 O2 Loeblich and Tappan, 1961

32 DISCUSSION.—See Olsson and others (1999:45).

AO1

P18 O1 Genus Acarinina Subbotina, 1953

33
P17
Truncorotaloides Brönnimann and Bermúdez, 1953.
34 AE10 Pseudogloboquadrina Jenkins, 1965b:1122.
E16
EOCENE

Muricoglobigerina Blow, 1979:1118.

LATE

P16 E15
AE9

35 TYPE SPECIES.—Acarinina acarinata Subbotina,

1953 (= junior subjective synonym of Globigerina nitida
FIGURE 13.1. Stratigraphic ranges of Oligocene species of Acarin- Martin, 1943; see Olsson and others, 1999).
ina discussed in this chapter. BKSA, 1995 = Berggren and others,
1995; K&S, 1983 = Kennett and Srinivasan, 1983; WPBP, 2011 =
Wade and others, 2011.
DISCUSSION.—See Olsson and others (1999:46) and
Berggren and others (2006:261) for general comments.

394

Acarinina collactea (Finlay, 1939)
Plate 13.1, Figures 1-16
(Pl. 13.1, Figs. 1-3: re-illustration of holotype of
Globorotalia collactea Finlay)
See Berggren and others (2006:276) for full synonymy list.
Globorotalia collactea Finlay, 1939:327, pl. 29, figs. 164,
165 [middle Eocene, Hampden Beach, North Otago,
New Zealand].
Truncorotaloides collactea (Finlay).—Jenkins, 1965a:843-
848, figs. 1-27 [middle Eocene, Hampden Beach, North
Otago, New Zealand].
Acarinina collactea (Finlay).—Pearson and Wade, 2015:
25, fig. 30.1 [lower Oligocene Zone O1, TDP Site 11,
Tanzania].
DESCRIPTION.
Type of wall: Moderately to coarsely muricate,
normal perforate, nonspinose.
Test morphology: Chambers arranged in a low
to moderate trochospiral, test compact, typically 4½
-5 chambers in the final whorl, slowly increasing in
size; peripheral outline weakly lobate; chambers on the
umbilical side wedge-shaped or triangular with blunt
and blocky muricae; sutures distinct, straight, radial and
wide; umbilicus narrow, deep; umbilical-extraumbilical
aperture; weakly convex to flat; on spiral side 10-12
sub-circular chambers in three whorls; sutures weakly
incised, curved; rounded to subangular peripheral
margin in edge view (modified after Berggren and others,
2006).
Size: Maximum diameter of holotype 0.18 mm,
thickness 0.13 mm.
DISTINGUISHING FEATURES.— Acarinina collactea
is characterized by its compact test, subangular chambers
that increase slowly in size, and wide, incised sutures.
It differs from Acarinina medizzai and A. echinata by
its lower trochospire, wedge shaped and more angular
chambers.
DISCUSSION.— Berggren and others (2006) noted
that this taxon is a particularly common form in mid-
high latitude assemblages, but the extension of the
range of Acarinina collactea into the upper Eocene
was controversial. Acarinina collactea is the most
Chapter 13 - Acarinina
common Oligocene acarininid. Here we extend the
range of this taxon further to the upper Oligocene (Zone
O7). Berggren and others (2006) restricted Acarinina
collactea to exclusively dextrally coiled specimens (see
their plate 9.8). We illustrate specimens with angular
chambers and wide sutures that are consistent with
Acarinina collactea although they are sinistrally coiled
(Plate 13.1, Figs. 11-16). Jenkins (1965a) found 95% of
specimens from the type locality to be dextrally coiled.
PHYLOGENETIC RELATIONSHIPS.— Berggren
and others (2006) tentatively suggested that Acarinina
collactea evolved from Acarinina pentacamerata.
Acarinina collactea probably gave rise to A. medizzai.
STRATIGRAPHIC RANGE.— Zone E7 (Berggren and
others, 2006) to Zone O7 (this study), possibly ranges
into the Miocene.
TYPE LEVEL.— Middle Eocene, Hampden Beach,
North Otago, New Zealand.
GEOGRAPHIC DISTRIBUTION.— Cosmopolitan
distribution in the Eocene. Our Oligocene records are
from the Adriatic Sea and western Indian Ocean.
STABLE ISOTOPE PALEOBIOLOGY.— No data
available.
REPOSITORY.— Holotype (TF 1150/1) and paratypes
deposited in collections of the New Zealand Geological
Survey Survey Collection, Lower Hutt, New Zealand.
Acarinina echinata (Bolli, 1957)
Plate 13.2, Figures 1-16
(Pl. 13.2, Figs. 1-3: re-illustration of holotype of
Catapsydrax echinatus Bolli)
See Berggren and others (2006:284) for full synonymy
list and taxonomic discussion.
Catapsydrax echinatus Bolli, 1957:165, pl. 37, figs. 2-5
[middle Eocene Porticulasphaera mexicana Zone,
Navet Fm., Trinidad].Huber, 1991:439, pl. 5, figs.
17, 20 [lower Oligocene Zone AP13, ODP Hole 744A,
Kerguelen Plateau].
Acarinina sp. (Bolli).—Grimm and others, 2005:246, pl. 1,
fig. 11 [lower Oligocene, Bodenheim Fm., Mainz Basin,
Germany].
395
Wade and Hernitz Kucenjak

396 Plate 13.1 Acarinina collactea (Finlay, 1939)


DESCRIPTION.
Type of wall: Moderately to coarsely muricate,
normal perforate, nonspinose.
Test morphology: Overall outline circular to
subcircular. “Test compact, biconvex, peripheral outline
weakly to moderately lobate, axial periphery rounded,
more rarely becoming slightly subangular; chambers
globular, 11-13 in adult tests increasing rapidly in size
until ultimate chamber, which is usually kummerform
and often connected to a variably shaped bulla that often
covers the umbilicus and part or most of the umbilical
sutures, 3½-4 chambers in the final whorl; umbilical
sutures radial, weakly to moderately depressed, spiral
sutures radial, weakly depressed to indistinct; aperture
variable in size, shape, position and number depending
on the characteristics of the bullate final chamber,
usually a single low arched opening directed towards the
umbilicus and surrounded by a narrow lip, more rarely
two or three small, low arched openings directed along
umbilical sutures” (Berggren and others, 2006:284).
Size: Maximum diameter of holotype 0.30 mm,
thickness 0.25 mm.
DISTINGUISHING FEATURES.— “Characterized
by its compact coiling, muricate surface texture, and
presence of a kummerform or bullate final chamber
and/or sutural bullae. Differs from Acarinina medizzai
by lacking a visible umbilicus and presence of
kummerform or bullate final chamber” (Berggren and
others, 2006:284).
DISCUSSION.— Among the Oligocene acarininids
Acarinina echinata is not very common.
PHYLOGENETIC RELATIONSHIPS.— Berggren
and others (2006) suggested that Acarinina echinata
may have evolved from Acarinina pseudosubsphaerica
during the middle Eocene. There are no known
descendants.
STRATIGRAPHIC RANGE.— The range of Acarinina
echinata is poorly constrained. It was described from
Zone E10 (Bolli, 1957), our highest recorded specimens
Plate 13.1 Acarinina collactea (Finlay, 1939)
1-3, holotype (Berggren and others, 2006, pl. 9.8, figs. 1-3), middle Eocene, Hampden Beach, New Zealand; 4, Zone O4/O5, Istra
More-3 well, 1045 m, Adriatic Sea; 5-8, (5, Pearson and Wade, 2015, pl. 30, fig. 1a, b) Zone O1, TDP Site 11/19/1, 60-75 cm, Stakishari,
Tanzania; 9-10, 13-14, Zone O1, DSDP Site 242, 10/5, 100-103 cm, western Indian Ocean; 11-12, Zone E16, Istra More-3 well, 1295 m,
Adriatic Sea; 15-16, Zone O5, Istra More-3 well, 968-974 m, Adriatic Sea. Scale bar: 4, 9, 11, 13-16 = 50 µm; 1-3 = 40 µm; 5-7, 12 = 20
µm; 8, 10 = 10 µm.
Chapter 13 - Acarinina
are from Zone O3/O4.
TYPE LEVEL.— Middle Eocene Porticulasphaera
mexicana Zone, Navet Fm., Trinidad.
GEOGRAPHIC DISTRIBUTION.— Rare, but globally
distributed in high and low latitudes. We have recorded
Oligocene occurrences from the Indian Ocean, Pacific
Ocean, Paratethys, and Kerguelen Plateau.
STABLE ISOTOPE PALEOBIOLOGY.— No data
available.
REPOSITORY.— Holotype (USNM P5729) deposited
at the Smithsonian Museum of Natural History,
Washington, D.C.
Acarinina medizzai (Toumarkine and Bolli, 1975)
Plate 13.3, Figures 1-16
(Pl. 13.3, Figs. 1-4: re-illustration of holotype and
paratype of Globigerina medizzai Toumarkine and
Bolli)
See Berggren and others (2006:294) for full synonymy
list and taxonomic discussion (and also discussion,
below).
Globigerina medizzai Toumarkine and Bolli, 1975:77, pl. 6,
figs. 1-8, pl. 5, figs. 8, 10, 13-15, 17, 19-22 [upper Eocene,
Scaglia Cinerea, Globigerinatheka semi-involuta Zone,
Possagno, Treviso, Northern Italy].
DESCRIPTION.
Type of wall: Moderately to coarsely muricate,
normal perforate, nonspinose.
Test morphology: Test relatively small,
biconvex, quadrate to subcircular in outline, weakly to
moderately lobate with a rounded periphery; chambers
globular, 8-12 arranged in a low trochospire of 3-4
whorls, increasing moderately in size, 4-6 in the final
whorl, rarely kummerform; umbilicus narrow in more
compact specimens, shallow and broad in more evolute
397
Wade and Hernitz Kucenjak

398 Plate 13.2 Acarinina echinata (Bolli, 1957)


forms; sutures weakly depressed, radial on both sides;
aperture a small and often indistinct low umbilical to
slightly extraumbilical arch (modified from Berggren
and others, 2006).
Size: Maximum diameter of holotype 0.19 mm;
test sizes range from 0.16 to 0.20 mm.
DISTINGUISHING FEATURES.— Acarinina medizzai
differs from A. collactea in its more rounded chambers,
lobate outline and more open umbilicus. It “differs from
A. echinata in the absence of a bulla and in the presence
of an umbilicus” (Berggren and others, 2006:294). It is
distinguished from Dipsidripella danvillensis (Howe
and Wallace) by its muricate wall texture and lower
arched aperture.
DISCUSSION.— Acarinina medizzai is not common in
the Oligocene. See discussion in Berggren and others
(2006).
PHYLOGENETIC RELATIONSHIPS.— Probably
descended from Acarinina collactea.
STRATIGRAPHIC RANGE.— The range is poorly
constrained. Acarinina medizzai evolved in middle
Eocene Zone E10 (Berggren and others, 2006). Our
youngest recorded occurrences are from Zone O4.
TYPE LEVEL.— Upper Eocene, Scaglia Cinerea,
Globigerinatheka semi-involuta Zone, Possagno,
Treviso, Northern Italy.
GEOGRAPHIC DISTRIBUTION.— In the Oligocene
we have recorded this species from the Adriatic Sea and
western Indian Ocean.
STABLE ISOTOPIC PALEOBIOLOGY.— No data
available.
REPOSITORY.— Holotype (C 24315) and paratypes
(C 27327-27329) deposited at the Basel Museum of
Natural History.
Plate 13.2 Acarinina echinata (Bolli, 1957)
1-3 (holotype, USNM 5729), Zone E12, Navet Fm., Trinidad; 4 (Grimm and others, 2005, pl. 1, fig. 11), lower Oligocene, Bodenheim
Fm., Mainz Basin, Germany; 5-6, 15-16, Zone O1, DSDP Site 242, 10/4, 100-103 cm, western Indian Ocean; 7-8, Zone O2, DSDP Hole
516F, 30/6 125-126 cm, Rio Grande Rise, South Atlantic Ocean; 9-11, 12 (9-11, same specimen), (Berggren and others, 2006, pl. 9.10,
figs. 6-8, 22), Zone AO1, ODP Hole 744A/15H/CC, Kerguelen Plateau, South Indian Ocean; 13-14, Zone O1, DSDP Site 242, 10/6, 106-
109 cm, western Indian Ocean. Scale bar: 1-5, 7, 9-12, 15 = 100 µm; 13 = 50 µm; 6, 8, 14, 16 = 10 µm.
Chapter 13 - Acarinina
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400 Plate 13.3 Acarinina medizzai (Toumarkine and Bolli, 1975)

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radiation of Paleocene planktic foraminifera: Paleobiology, v. biostratigraphy, and phylogeny of Oligocene Acarinina,
22, p. 461-480. in Wade, B.S., Olsson, R.K., Pearson, P.N., Huber, B.T.
Olsson, R.K., Hemleben, Ch., Berggren, W.A., and Huber, B.T.,
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planctoniques de l’Eocene moyen et supérieur de la Coupe de

Plate 13.3 Acarinina medizzai (Toumarkine and Bolli, 1975)

1-4 (1-3, holotype, Toumarkine and Bolli, 1975, pl. 6, figs. 1, 7, 8; 4, paratype, Toumarkine and Bolli, 1975, pl. 6, fig. 3), Zone E14,
sample 68/60, Italy; 5-6, Zone O2, Istra More-3 well, 1175 m, Adriatic Sea; 7-11, Zone O4, Istra More-4 well, 1225-1231 m, Adriatic
Sea; 12, Zone O1, Istra More-3 well, 1270 m, Adriatic Sea; 13-14, Zone O1, TDP Site 17/22/1, 0-15 cm, Stakishari, Tanzania; 15-16,
Zone O4, Istra More-4 well, 1225-1231 m, Adriatic Sea. Scale bar: 13 = 100 µm; 1, 4-5, 7, 9, 11-12, 15 = 50 µm; 6, 8, 10, 16 = 20 µm; 2,
14 = 10 μm; 3 = 5 µm.

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Wade and Hernitz Kucenjak

402